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Genetic Analysis of Needle Morphological and Anatomical Traits Among Nature Populations of Pinus Tabuliformis

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Genetic Analysis of Needle Morphological and Anatomical Traits Among Nature Populations of Pinus Tabuliformis Journal of Plant Studies; Vol. 6, No. 1; 2017 ISSN 1927-0461 E-ISSN 1927-047X Published by Canadian Center of Science and Education Genetic Analysis of Needle Morphological and Anatomical Traits among Nature Populations of Pinus Tabuliformis Mei Zhang1, Jing-Xiang Meng1, Zi-Jie Zhang1, Song-Lin Zhu2 & Yue Li1 1National Engineering Laboratory for Forest Tree Breeding, Key Laboratory for Genetics and Breeding of Forest Trees and Ornamental Plants of Ministry of Education, College of Biological Sciences and Technology, Beijing Forestry University, Beijing 100083, China 2The Forestry Bureau of Xixian, China Correspondence: Mei Zhang, College of Biological Sciences and Technology, Beijing Forestry University, Beijing 100083, China. E-mail: [email protected] Received: December 6, 2016 Accepted: January 10, 2017 Online Published: January 21, 2017 doi:10.5539/jps.v6n1p62 URL: http://dx.doi.org/10.5539/jps.v6n1p62 Abstract The morphological and anatomical traits of needles are important to evaluate geographic variation and population dynamics of conifer species. Variations of morphological and anatomical needle traits in coniferous species are considered to be the consequence of genetic evolution, and be used in geographic variation and ecological studies, etc. Pinus tabuliformis is a particular native coniferous species in northern and central China. For understanding its adaptive evolution in needle traits, the needle samplings of 10 geographic populations were collected from a 30yr provenience common garden trail that might eliminate site environment effect and show genetic variation among populations and 20 needle morphological and anatomical traits were involved. The results showed that variations among and within populations were significantly different over all the measured traits and the variance components within population were generally higher than that among populations in the most measured needle traits. Population heritabilities in all measured traits were higher than 0.7 in common garden sampling among populations. Needle traits were more significantly correlated with longitude than other factors. First five principal components accounted for 81.6% of the variation with eigenvalues greater than 1; the differences among populations were mainly dependent on needle width, stomatal density, section areas of vascular bundle, total resin canals, and mesophyll, as well as area ratio traits. Ten populations were divided into two categories by Euclidean distance. Variations in needle traits among the populations have shown systematic microevolution in terms of geographic impact on P. tabuliformis. This study would provide empirical data to characterize adaptation and genetic variation of P. tabuliformis, which should be more available for ecological studies. Keywords: genetic structure, needle, morphological and anatomical traits, geographic population, Pinus tabuliformis 1. Introduction Pinus tabuliformis is a native conifer species in northern and central China. It survived from the quaternary glacial with complex genetic components and climate factors widely changed in its distributions (Hewitt, 2000; Chen, 2007; Guo, 2008). Significantly variations among nature populations in P. tabuliformis have been reported on growth, wood property, physiology and propagation traits (Mao et al., 2009, Mao et al., 2011; Xu et al., 1991; Yuan et al., 2014; Niu et al., 2013; Yang et al., 2015). Evidences from molecular have indicated the genetic differentiation among populations bounded up with the geographic distance (Chen, 2007; Wang, 2010; Gao, 2009). Like other plant in temperate arid steppe zone, P. tabuliformis have formed phenotypic characters to resist lower temperature, water depletion and light conditions (Zhang, 2010; Liu, 2012). Needle is the most vigorous assimilation organs for conifer species, and the morphological and anatomical traits of needle are important references to plant taxonomy (Anna K. et al., 2013; Xing, 2014; Huang, 2016). Needle traits are closely associated with physiological and functional attributes of plants including photosynthesis, respiration, water metabolism, nutritional status, as well as stress resistance (Oleksyn et al., 1997; Eguchi et al., 2004; Wu, 2007; Mao et al., 2012). Phenotypic variations of needle morphological and anatomical characteristics were considered the results in physiological and adaptive genetic evolution (Anna K. et al., 2013; 62 http://jps.ccsenet.org Journal of Plant Studies Vol. 6, No. 1; 2017 Balkrishna et al., 2014), and have been widely used as available indicators in geographic variation, phylogenesis and evolutionary studies (Cole et al., 2007; Xing, 2014; Melville, 2002, B. Nikolić et al., 2013; Androsiuk Piotr et al., 2011; K. Boratynska et al., 2009). However, phenotypic variations of needle traits should be influenced by both environment and genetic impacts (Li, 2009; Xu,1991; Cole et al., 2008) in studies with samples from natural forests within species (Legoshchina et al., 2013; Michael et al., 2012), hardly tell the amount of genetic or environment contributions and limit in explaining the real effects on changes among populations. Variations of needle traits from population samples based on common garden trail could provide the genetic contributions among populations. The research and analysis of P. tabuliformis mainly involve the natural distribution characters of needles, and results that might be limited because of environmental variations at different population sites (Nikolić et al., 2013; Xing et al., 2014). In this study, we collected the needle samples from a 30yr provenience common garden trail with ten populations in P. tabuliformis in order to eliminate environment effects on needle traits among populations. The objectives of this study were to (1) reveal the variations in needle morphological and anatomical traits among populations; (2) illustrate the variations in needle traits among individuals within population; (3) evaluate the genetic and environment impacts on phenotypic variation for each needle trait; and (4) clarify the phylogenetic relationships among populations. This study provides theoretical and methodological reference for the conservation biology of populations in the coniferous morphological variation, ecological adaptability, system evolution and population genetics. Provide the basis for the use of coniferous morphological traits in each study area. 2. Materials and Methods 2.1 Sample Populations Ten geographic populations, located in typical habitat of P. tabuliformis, were involved (Fig. 1) in our experiment. Ten populations are as follows: Heilihe, Inner Mongolia(HLH); Dongling, Heibei(DL); Lingkongshan, Shanxi(LKS); Guandishan, Shanxi(GDS); Shangzhuang, Shanxi(SZ); Nanyang, Henan(NY); Luonan, Shannxi(LN); Shuanglong, Shannxi(SL); Xiaolongshan, Gansu(XLS); Huzhu, Qinghai(HZ). Figure 1. Location of ten P. tabuliformis populations sampled in North China The environmental information of sampled populations is shown in Table 1. The annual mean temperatures and precipitations in each geographic location of populations were obtained from www.worldclim.com (Hijmans et al., 2005). 63 http://jps.ccsenet.org Journal of Plant Studies Vol. 6, No. 1; 2017 Table 1. Sampling populations of P. tabuliformis Population Sample size Longitude(°E) Latitude (°N ) Altitude(m) Annual mean temperature(°C) Annual precipitation (mm) HZ 29 102°28′ 36°58′ 2300 2.4 443 XLS 21 106°00′ 34°20′ 1630 8.5 663 NY 14 112°03′ 33°32′ 810 12.6 776 LNGC 12 110°21′ 34°21′ 1220 10.0 720 HLH 25 118°58′ 42°17′ 1300 7.6 360 GDS 19 111°29′ 37°54′ 1500 1.5 561 SL 25 108°56′ 35°41′ 1650 10.2 543 LKS 13 112°02′ 36°37′ 1665 5.6 604 SZ 13 111°12′ 36°46′ 1660 6.2 545 DL 14 117°38′ 40°11′ 200 10.6 587 2.2 Field Experimental Design The common garden test of proveniences in P. tabuliformis was conducted in Xixian, Shanxi province (111°10′E, 36°48′N), the distribution central of the species and represented the general environmental conditions of the species. The experiment was established in 1981 with 2a seedlings from ten nature populations from the central to the edge of geographic distribution regions. A Randomized Complete Block Design (RCBD) with 6 blocks (replication), 24-individuals rectangular plot was set for the trail, 2m×2m in planting space. However, the remaining plants for each population were varied after 30yr nature selection. Needle samples from the provenience test were collected in September on 30yr plants in 2011, 11-30 individual trees for each population were involved from randomized sample trees. For each plant, 3 needles would be randomly selected for analysis. We examined a total of 20 traits involving morphology, anatomy and ratio of needle structures (Table 2). All the traits were adaptive- related and have been reported in previous researches (Huang et al., 2016; Xing et al., 2014; Zhao et al., 2008). 2.3 Measurement of Needle Traits Twenty needle traits were measured according to Xing’s work (Xing et al., 2014), including 16 directly measured morphological and anatomical traits and four area-ratio indices (the ratios between needle organizational structures in needle cross sections) (Table 2). (1) Morphological traits (Fig. 2A): seven morphological traits were measured, including needle length (NL, 1 cm); the width and thickness in the middle part of the needle (NW and NT, 0.1 mm); the number of stomatal rows on the front and back sides of the needle (CSRN and FSRN, measured by stereomicroscope);the number of stomata rows(NSR, calculated as NSR=( FSRN+CSRN));
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